Vestnik zoologii, 51(6): 453–470, 2017 Fauna and Systematics DOI 10.2478/vzoo-2017-0056

UDC 595.773.4(4:479:5–15) THE CARPOMYINI FRUIT (DIPTERA: ) OF , CAUCASUS, AND MIDDLE EAST: NEW RECORDS OF PESTS, WITH IMPROVED KEYS

V. A. Korneyev1, 4, R. I. Mishustin2, S. V. Korneyev3

1, 3Schmalhausen Institute of Zoology, NAS of , vul. B. Khmelnytskogo, 15, Kyiv, 01030 Ukraine 1E-mail: [email protected] 2UkrainianEntomological Society E-mail: [email protected] 3E-mail: [email protected] 4Corresponding author

Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus, and Middle East: New Records of Pest Species, with Improved Keys. Korneyev, V. A., Mishustin, R. I., Korneyev, S. V. — Based on previously unpublished specimens from the collection of I. I. Schmalhausen Institute of Zoology (Kyiv), all the species of the tribe Carpomyini known to occur in Europe and Asia westwards of the Caspian Sea and Afghanistan, are listed. Th e melon fruit fl y, Carrpomya (Myiopardalis) pardalina (Bigot) is recorded for the fi rst time from Ukraine and Europe, and (s. str.) vesuviana (A. Costa, 1854) for continental Ukraine, based on collection material. Carpomya (Goniglossum) liat (Freidberg, 2016), comb. n., is established as Goniglossum has not been adopted at the rank. New records and illustrated keys to species are given. Key words: Diptera, Tephritidae, Carpomya, , Europe, Caucasus, Israel, Turkey, Iran.

Introduction

Th e fruit fl ies of the tribe Carpomyini occur predominantly in the New World, (Korneyev, 1996; Smith & Bush, 1999) with 35 species occurring in the Palaearctic Region, and six or more species in the Oriental Region (Nepal, India, Th ailand, Cambodia). Two pest species, and R. completa, have been introduced from North America to Europe (Merz, 1991) and two others, Carpomya pardalina and C. vesuviana, — from the Middle East into the Oriental Region (Hardy, 1973, 1977). Some species are serious pests of fruits (including cherries, melons, walnuts, and apples) as well as wild berries (Cornus, Berberis, Rosa, etc.) and are subjects for quarantine control. 454 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Th e Palaearctic species of Rhagoletis (including Zonosema, Megarrhagoletis, and Microrhagoletis now considered its synonyms) were reviewed and keyed by Rohdendorf (1961). Later, larval morphology of known and some additional species was described by Kandybina (1961, 1962, 1972, 1977), Kandybina & Richter (1976), and Richter & Kandybina (1997). Th e Middle Asian, Far East, and Oriental species of Carpomya and Rhagoletis were keyed by Korneyev & Merz (1997), Korneyev & Ovchinnikova (2004), and Ito (2011). Freidberg & Kugler (1989), Merz (1994), Korneyev (1997), and Mohamadzade Namin & Rasoulian (2009) provided keys to the European, Middle Eastern, and Caucasian species, but no keys covering the whole Western Palaearctics existed. In this paper, keys to all the species of Europe, Asia Minor, Caucasus, and Near East including Iran, are provided, along with references to each species, data on its known host plants and distribution, as well as some taxonomic remarks are provided.

Material Th e specimens mentioned in this paper are deposited mostly in the collection of SIZK — Schmalhausen Institute of Zoology, Kyiv, Ukraine; except a few specimens from MHNG — Muséum d’Histoire Naturelle, Genève.

Carpomya Costa, 1854 Carpomya Costa, 1854: 87. Type species: Carpomya vesuviana Costa, 1854 (by monotypy). Goniglossum Rondani, 1856: 110. Type species: wiedemanni Meigen (by original designation). Myiopardalis Bezzi 1910: 10 Type species: Carpomyia pardalina Bigot (by original designation). Carpomyia Rondani, 1870: 22, misspelling of Carpomya Costa. Gonyglossum Effl atoun, 1924: 19, misspelling of Goniglossum Rondani. Gonioglossum Hendel, 1914: 90, misspelling of Goniglossum Rondani.

Diagnosis. Middle-sized (3.5–5.0 mm) fruit fl ies with 3 frontal and 2 orbital setae, pale postocellar seta, variable shape of head, antenna and proboscis (short with fl at, slightly receding or vertical face, apically pointed fl agellomere 1 and short labellum, as on fi gs 2, 2–4, in Carpomya s. str. and Myiopardalis and long, with carinate, anteriorly produced face and long, geniculate proboscis as on fi g. 2, 1, in Goniglossum), usually brightly patterned, pale yellow to orange mesonotum with shiny black spots and grey microtrichose areas (if, in C. incompleta uniformly pale reddish yellow and microtrichose, then fl agellomere 1 pointed and ocellar setae short), long and strongly acute posterior lobe of surstylus of male, oviscape with T-shaped desclerotized posteromedial area ventrally, and aculeus either uniformly tapered apically (in Carpomya s. str. and Myiopardalis) or abruptly cut into a wide, serrate apex in Goniglossum (see Freidberg, 2016). Th ird instar larva with a few (3–4) serrate oral ridges and stomal sensory organ with strong preoral teeth. Remarks. Carpomya shows no signifi cant diff erences from the largest genus of Carpomyini, Rhagoletis, except for the mesonotal pattern as on fi gs 1, 1, 3, 1–3, 3 and 3, 5 (though C. incompleta has uniformly pale reddish yellow mesonotum, as in some Rhagoletis and can be distinguished from them only by having short ocellar seta, which is long in all Rhagoletis) species. Neither Carpomya nor Rhagoletis have been defi nitively proven to be monophyletic (V. Korneyev, unpublished data; J. Jenkins, unpublished data; J. Smith et al., in prep.): Carpomya is either the sister-group of Rhagoletis, or an in-group within the species assigned to Rhagoletis. Taxonomic consequences and concepts of both genera likely will be infl uenced by the results of forthcoming phylogenetic analyses based on the sequences of multiple genes (J. Smith et al., in prep.), and we abstain from any taxonomic changes (including those by Freidberg, 2016) until their phylogenetic relationships are more clearly resolved. Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 455

Fig. 1. Carpomyini. 1–2 — Carpomya pardalina, Kherson (1 — freshly emerged male; 2 — infested melon with holes in skin (cyan arrows) prepared by larvae for emerging of adults); 3 — Rhagoletis meigenii on Berberis fruit, Berlin; 4 — R. cerasi on Lonicera fruit, Kyiv, Grishko Botanical Gardens. Photos by R. I. Mishustin (1–2) and V. A. Korneyev (3–4).

Key to species of Carpomya of the Old World 1. Proboscis with extremely long and narrow labellum (fi g. 2, 1). Head shape: face with ventral margin produced anteriorly and strongly developed medial carina (fi g. 2, 1). Cell bcu closed by arcuate vein without lobe (fi g. 4, 1). Flagellomere 1 apically rounded. Scutellum with large medial black spot on disk (fi g. 3, 1–2). In fruits of Bryonia...... C. (Goniglossum) 2. — Proboscis with fl eshy and moderately short labellum (fi gs 2, 2–4). Head shape: face with slightly pro- duced carina, ventral margin not strongly produced anteriorly (fi gs 2, 2–4). Cell bcu with posteroapical lobe (fi gs 4, 2–5). Flagellomere 1 apically pointed. Scutellum either entirely yellow (fi g. 3, 4), or only with marginal spots (fi g. 3, 5), or at most with small medial black dot on disk (fi g. 3, 3)...... 3. 2. Medial black spot on scutellum disk very large, confl uent with black dots at bases of apical scutellar setae (fi g. 3, 1). In fruits of Bryonia cretica L. and B. syriaca Boiss. Southern Turkey, Israel...... C. (Goniglossum) liat (Freidberg( — Medial black spot on scutellum disk smaller, separate from black dots at bases of apical scutellar setae (fi g. 3, 2). In fruits of L. and B. dioica Jacq. Europe, Caucasus...... C. (Goniglossum) wiedemanni (Meigen) 3. Face high, longer than frons and 3 times as long as fl agellomere 1 (fi g. 2, 2). Scutellum with small medial black dot on its disk (fi g. 3, 3). In fruits of melons...... C. (Myopardalis) pardalina Bigot — Face shorter, at most 2 times as long as fl agellomere 1 (fi gs 2, 3–4). Scutellum without medial black dot on its disk (fi g. 3, 4–5). Not in melons ...... C. (Carpomya s. str.) 3. 4. Apical crossband absent (fi g. 4, 3). Scutum and scutellum entirely reddish yellow (fi g. 3, 4). In fruits of Ziziphus...... C. incompleta (Becker) — Wing with 4 yellow crossbands (fi gs 4, 4–5). Scutum with dark pattern; scutellum with pair of large black spots at posterior margin (fi g. 3, 5)...... 4. 5. Anepisternum and mediotergite (below scutellum) yellow. Ocellar seta setula-like, almost inconspicu- ous. In Ziziphus...... C. vesuviana A. Costa — Anepisternum and mediotergite black. Ocellar seta usually as long as orbital setae. In Rosa fruits...... C. schineri Loew 456 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Carpomya (Carpomya) incompleta (Becker, 1903) (fi gs 2, 3; 3, 4; 4, 3)

Trypeta incompleta Becker, 1903: 135; Carpomyia incompleta: Bezzi, 1910: 11; Silvestri, 1916: 179; Effl atoun, 1924: 40; Hendel, 1927: 92; Carpomya incompleta: White & Elson-Harris, 1992: 281; Norrbom, 1997: 340; Norrbom et al., 1999: 115; Korneyev & Dirlbek, 2001: 479.

Material. Israel: Ein Gedi, –400 m, swept from Ziziphus, 31.05.2000, 3 {, 2 } (V. Korneyev) (SIZK).

Distribution. ; Egypt, Sudan, Eritrea; Israel, Iraq. Host plants. Ziziphus jujuba Mill., Z. lotus L., Z. spina-christi (L.) Willd., Z. sativus Gaertn. (Rhamnaceae) (Freidberg & Kugler, 1989; Smith & Bush, 1999).

Carpomya (Carpomya) schineri (Loew, 1856) (fi gs 2, 4; 3, 5; 4, 4)

Trypeta schineri Loew, 1856: 52; schineri: Loew, 1862b: 49; Myiopardalis schineri: Bezzi, 1910: 9; Carpomyia schineri: Hendel, 1927: 92; Kandybina, 1965: 668, 1977: 117; Korneyev, 1983: 12; Kameneva & Korneyev, 1985: 68; Smit, 2010: 127; Carpomya schineri: White & Elson-Harris, 1992: 285; Norrbom, 1997: 340; Norrbom et al., 1999: 115.

Fig. 2. Carpomya spp. heads, left lateral view: 1 — C. (Goniglossum) liat; 2 — C. (Myiopardalis) pardalina; 3 — C. (s. str.) incompleta; 4 — C. (s. str.) schineri. Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 457

Fig. 3. Carpomya spp. mesonotums, dorsal view: 1 — C. (Goniglossum) wiedemanni; 2 — C. (G.) liat; 3 — C. (Myiopardalis) pardalina; 4 — C. (s. str.) incompleta; 5 — C. (s. str.) schineri.

Fig. 4. Carpomya spp. wings: 1 — C. (Goniglossum) wiedemanni; 2 — C. (Myiopardalis) pardalina; 3 — C. (s. str.) incompleta; 4 — C. (s. str.) schineri; 5 — C. (s. str.) vesuviana. 458 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Material. Georgia: [no locality], 5.10.1959, ex fruits of Rosa sp. 3 {, 1 } [no collector name]; Ukraine: Cherkasy: Kaniv, 24.07.1957, 1 { [no collector name]; Mykolaiv: Trykraty, Aktove, 47.70 N 31.43 E, 1 {, 1 } (S. & V. Korneyev); Donetsk: Kamyani Mohyly Nature Reserve, ex Rosa fruit, 10.1992–04.1993, 1 } (V. Korneyev); Crimea: Karadagh, 07.1984, 2 { (Karachevskaya) (SIZK).

Distribution. , , Bulgaria, European Russia (Centre, East, and South), (mainland), , , Italy (mainland), , , (mainland), , Ukraine; Asian Russia (Northern Caucasus; East Siberia, Far East), Georgia, Israel, Turkey, Kazakhstan, Kyrgyzstan. Host plants. Rosa canina L., R. beggerana Schrenk, R. damascena Mill., R. gallica L. R. kokanica (Rgl.) Juz., R. pulverulenta M.B., R. rubiginosa L., R. rugosa Th unb., R. spinosissima L., R. villosa L., (Rosaceae) (Hendel, 1927; Freidberg & Kugler, 1989; Kandybina, 1977; Smith & Bush, 1999).

Carpomya (Carpomya) vesuviana A.Costa, 1854 (fi g. 4, 5) Carpomya vesuviana Costa, 1854: 175; White & Elson-Harris, 1992: 283; Norrbom, 1997: 340; Norrbom et al., 1999: 115; Mohamadzade Namin, 2016; Carpomyia vesuviana: Bezzi, 1910: 10; Silvestri, 1916: 179; Hendel, 1927: 93; Kandybina, 1965: 666, 1977: 112; Trikoz & Litvinova, 2007: 74; Mamedov et al., 2015. Material. Ukraine: Kherson, in fruits of Ziziphus, 07.2016, 4 third-instar larvae (R. Mishustin) (SIZK).

Distribution. Italy, Moldova (unconfi rmed records), Ukraine (fi rst confi rmed record); Asian Russia (North Caucasus), Afghanistan, Azerbaijan, Cyprus, Iran, Iraq, Is- rael, Lebanon, Syria, Tajikistan, Turkey, Turkmenistan, Uzbekistan; Cambodia, India, Pa- kistan, Th ailand; Mauritius. Host plants. Ziziphus jujuba Mill., Z. mauritiana Lam., Z. nummularia (Burm.) Wight and Arn., Z. rotundifolius Lam., Z. sativus Gaertn. (Rhamnaceae) (Freidberg & Ku- gler, 1989; Smith & Bush, 1999). Remarks. Trikoz & Litvinova (2007) and various Internet sites report C. vesuviana from Crimea (since 1998) and Kherson Region in Ukraine, but no material in collections was available for identifi cation. Now this species is considered to be well established pest in all the areas of planting of Ziziphus in Southern Ukraine.

Carpomya (Goniglossum) liat (Freidberg, 2016), comb. n. (fi gs 2, 1, 3, 2) : Freidberg & Kugler, 1989: 189 (misidentifi cation). — Goniglossum liat Freidberg, 2016: 59.

Material. Type: Paratypes 3 {, 4 }: Israel: Nizzanim, 03–04.05.2005, 3 {, 2 } (A. Freidberg) (SIZK). Non-type: Israel: [Nizzanim], ex fruits of Bryonia sp., coll. 1.05 — exit 30.05.2000, 2 {, 1 } (A. Freidberg) (SIZK).

Distribution. Israel, Turkey (South). Host plants. Bryonia cretica L., B. syriaca Boiss. () (Freidberg & Ku- gler, 1989; Freidberg, 2016).

Remarks. Th is species was recently recognized and described by Freidberg (2016); its distribution apparently does not exceed the one mapped in that paper. As we consider Goniglossum (and Myiopardalis) as subgenera within Carpomya, G. liat is also transferred to the latter genus.

Carpomya (Goniglossum) wiedemanni (Meigen, 1826) (fi gs 3, 1; 4, 1) Trypeta wiedemanni Meigen, 1826: 320; Oedaspis wiedemanni: Loew, 1862 b: 49; Carpomya wiedemanni: Norrbom, 1997: 340; Norrbom et al., 1999: 115; Gonioglossum wiedemanni: Hendel, 1927: 92; Richter, 1960: 804; Kandybina, 1977: 112; Korneyev, 1983: 12; Kameneva & Korneyev, 1 985: 68; Goniglossum wiedemanni: Smit, 2010: 128; Freidberg, 2016: 59; Gonyglossum wiedemanni: Bezzi, 1910: 4, 6; Silvestri, 1920: 209. — Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 459 bryoniae Meigen, 1826: 321 (nomen nudum); Goniglossum wiedmanni: Rondani, 1856: 110 (misspelling of wiedemanni Meigen). Material. France: Gard: Pont du Gard, 43.95 N, 4.53 E, 5.06.1995, 1 {, 1 } (B. Merz & M. Eggenberger); Italy: Bognocavallo 10 km W Ravenna, Rt S253, 44.40 N, 12.05 E, ex , exit 10.04.2007, 3 {, 2 } (G. Pezzi); Ukraine: Crimea: Kerch Peninsula, Opuk Mt., 45.03 N, 36.22 E, slopes, 21.06.1950 (I. Maltsev); Armenia: Gudemnis, 38.95 N, 46.18 E, 22.06.1984, 4 { (V. Ermolenko) (SIZK).

Distribution. Austria, Belgium, Britain, European Russia (south); France (mainland and Corsica), Germany, Hungary, Italy (mainland and Sicily), Spain (mainland), Switzer- land, the , Ukraine; Armenia (fi rst record). Host plants. Bryonia dioica (Cucurbitaceae) (Merz, 1994); Bryonia cretica L., B. syri- aca Boiss. (Smith & Bush, 1999). Remarks. Th is species was recorded from Ukraine (Richter, 1960) based on speci- mens collected by Jaroszewsky in Myrhorod in the 19th century; the specimen from Crimea was collected almost 70 years ago. Th is species has not been collected in Ukraine recently, to our knowledge. Carpomya (Myopardalis) pardalina Bigot, 1891 (fi gs 1, 1–2; 2, 2; 3, 3; 4, 2) Carpomyia pardalina Bigot, 1891: 51; Myiopardalis pardalina: Bezzi 1910: 4, 8; Hendel, 1927: 90; Freidberg & Kugler, 1989: 194; White & Elson-Harris, 1992: 349; Carpomyia (Myiopardalis) pardalina: Zaitzev, 1919: 66; 1947: 6; Rohdendorf, 1936: 22, 35; Kandybina, 1965: 668; 1977: 51; Carpomya pardalina: Norrbom, 1997: 340; Norrbom et al., 1999: 115. — Carpomyia caucasica Zaitzev, 1919: 64; Carpomyia (Myiopardalis) caucasica: Rekatch, 1930: 5. — Kryshtal, 1949: 203 (misidentifi cation).

Material. Israel: Genin, 1.07.1971, 1 {, 1 } (A. Freidberg); Ukraine: Kherson: Skadovsk District, emerged from Cucumis melo, 17.09.1916, 1 { (R. Mishustin) (SIZK). Distribution. Ukraine (fi rst record in Europe); Asian Russia (North Caucasus); Cyprus, Iran, Iraq, Israel, Lebanon, Syria, Turkey, Armenia, Georgia, Azerbaijan, Turk- menistan, Afghanistan; India. Host plants. Cucurbitaceae: Cucumis melo, C. melo var. fl exuosus (Freidberg & Kugler, 1989). Remarks. It is believed that the melon fruit fl y is now widespread in all the areas of melon plantings within Ukraine, as it was entirely neglected at the beginning of its spread. Furthermore, it has been found to be imported into many other regions of Ukraine with melonstransported for commercial purposes; for instance, in the Transcarpatian Region (http://izan.kiev.ua/ukrbin/show_image.php?imageid = 41666). Its establishment, however, has been restricted to the southern Ukraine, which is the only appropriate area for cultivation of its hosts.

Rhagoletis Loew, 1862 Rhagoletis Loew, 1862b: 44 Type species: Rhagoletis cerasi (Linnaeus, 1758) (by monotypy). Zonosema Loew 1862b: 43 Type species: Tephritis alternata Fallén, by subsequent designation of Rondani, 1870: 6. Microrrhagoletis Rohdendorf, 1961: 187 Type species: Microrrhagoletis samojlovitshae Rohdendorf (by original designation). Megarrhagoletis Rohdendorf, 1961: 196. Type species: Megarrhagoletis magniterebra Rohdendorf (by original designation). Diagnosis. Medium-sized (3.0–8.0 mm) fruit fl ies with 3 frontal and 2 orbital setae, pale or dark postocellar seta, short head, pointed apex of fl agellomere 1 (except in R. kurentsovi (Rohdendorf, 1961) from Far East Russia and some Neotropical species); either pale yellow to orange, or mostly black with creamy white or yellow scutellum except 460 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Fig. 5. Rhagoletis spp. mesonotums (1–8), dorsal view, and head (2), lateral view: 1 — R. completa; 2 — R. bagheera; 3 — R. berberidis; 4 — R. cerasi; 5 — R. fl avicincta; 6 — R. fl avigenualis; 7 — R. obsoleta, Myhiia; 8 — R. sp. near obsoleta, Mt. Hermon. base and postpronotal lobes, long and variously acute posterior lobe of surstylus of male, oviscape with T-shaped desclerotized posteromedial area ventrally, and aculeus uniformly tapered apically. Th ird instar larva with variable number (from 3 to 20) of oral ridges and stomal sensory organ with or without preoral teeth.

Remarks. Rhagoletis is a heterogeneous genus with fi ve to seven groups of species associated with diff erent host plant families (Rhamnaceae, Berberidacea, Solanaceae, etc.) in the Palaearctic, Nearctic, and Neotropical Regions (and a few species in montane areas in the northern Oriental Region), with the relationships among them and the other genera of the tribe Carpomyini (e. g., Carpomya and Zonosemata) still poorly resolved and understood. In the case Carpomya is found to be an in-group within Rhagoletis, the latter either would incorporate its species and become a junior synonym of the latter, or should be split into several monophyletic genera (V. Korneyev, unpublished data; J. Jenkins, Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 461 unpublished data; J. Smith et al., in prep.). Here, we abstain from any taxonomic changes until phylogenetic relationships in Rhagoletis are better understood.

Key to species of Rhagoletis of western Palearctic Region 1. Mesonotum uniformly reddish-yellow (as on fi g. 5, 1), rarely brown posteriorly...... 2. — Scutum black; postpronotal lobe, sides and scutellum yellow or whitish (as on fi gs 5, 2–8)...... 5.

2. Wing with short accessory crossband in cell r1 (fi gs 6, 1–3)...... 3.

— Wing without accessory crossband in cell r1 (fi g. 7, 3). Larvae in walnut (Juglans) husk ...... ` ...... R. completa Cresson

3. Subapical band not extending anterior of vein R2+3 (usually not beyond R4+5) (fi g. 6, 1). Larvae in Berberis...... R. meigenii (Loew) — Subapical band reaching anterior wing margin (fi gs 6, 2–7)...... 4. 4. Abdominal tergites 1–5 entirely reddish-yellow. Larvae in Rosa...... R. alternata (Fallén) — Abdominal tergites 1–5 black basally or entirely. Larvae in Berberis. .R. caucasica Kandybina & Richter

5. Wing with short accessory crossband in cells r1 and r2+3 (fi gs 6, 4–6)...... 6.

— Wing without accessory crossband in cell r1 and r2+3 (fi gs 7, 1–8)...... 7. 6. Scutum with four silvery tomentose vittae; scutellum entirely yellow, except faint anterior band (fi g. 5, 4). Wing with apical crossband crossing vein M and entering into cell m (fi gs 6, 5–6). Larvae in Lonicera and cherry fruits...... R. cerasi (Linnaeus) — Scutum shining black; anterior margin of scutellum black (fi g. 5, 3). Wing with apical crossband touching M apex, but not entering into cell m (fi g. 6, 4). Larvae in Berberis seeds. .... R. berberidis Jermy 7. Apical crossband simple, not bifurcated (fi g. 7). [Fore femur colouration variable.] Larvae not in cherries. .. 8. — Apical crossband bifurcated posteriorly, usually with distal branch isolated as separate spot (fi g. 6, 7). [Fore femur usually entirely yellow.] Larvae in cherries ...... R. cingulata (Loew) 8. Apical crossband entirely contiguous with apical wing margin (fi gs 7, 1–2, 4)...... 9. — Apical crossband separated from apical wing margin by crescentic marginal hyaline area (fi gs 7, 5–8)...... 10. 9. Femora yellow. Larvae in Lonicera...... R. fl avicincta Enderlein — Femora black...... R. obsoleta Hering and R. sp. near obsoleta 10. Femora yellow. Larvae in Juniperus...... 11. — Femora black...... 12.

11. Discal and subapical crossbands widely connected at least in cell r2+3 (fi g. 7, 5)...... R. zernyi Hendel — Discal and subapical crossbands entirely separated (fi g. 7, 6)...... R. fl avigenualis Hering 12. Smaller: WL ♂ < 2.45mm (2.0–2.4mm), WL ♀ < 2.55mm (2.2–2.5 mm). Larvae in Rhamnus...... R. bagheera Richter & Kandybina — Larger: WL ♂ > 2.45mm (2.5–3.2mm), WL ♀ > 2.55mm (2.6–3.2mm)...... 13. 13. Fore femur mostly black, except yellowish ventral side. Larvae in Hippophae...... R. batava Hering — Fore femur mostly yellow, except brownish dorsal band. Larvae in Juniperus. .... R. sp. near fl avigenualis

Rhagoletis alternata (Fallén, 1814) (fi gs 6, 2) Tephritis alternata Fallén, 1814: 162, 1820: 3; Trypeta alternata Loew, 1844: 315; Zonosema alternata: Loew, 1862b: 43; Zonosema alternatum: Rohdendorf, 1961: 192; Rhagoletis alternata: Hendel, 1927: 74; Rohdendorf, 1936: 23; Bush, 1966: 512; Korneyev, 1983: 12; Kameneva & Korneyev, 1985: 68; White & Elson-Harris, 1992: 386; Merz, 1994: 108, 2001: 92; Korneyev, 1997: 95; Norrbom et al., 1999: 200; Merz & Korneyev, 2004; Smit, 2010: 129.

Material. Ukraine: Kyiv: “Kyiv”, 27.06.1920, 1 { [S. Paramonov], Irpin, 50.5279 N, 30.2643 E, ex Rosa sp., 7.09.1993–07.1994, 2 } (V. Korneyev); Chernihiv: Yaduty, 51.41 N, 32.39 E, ex Rosa sp., 08.2000– 1.06.2001, 1 {, 1 } (V. Korneyev); Cherkasy: Kaniv Nature Reserve, 49.70 N, 31.53 E, ex Rosa sp., 2.10.1980– 1.06.1981, 1 { (V. Korneyev); Russia: Northern Caucasus: Karachaevo-Cherkessia: Teberda, 43.44 N, 41.74 E, 18.07.2013, 1 {, 19.07.2013, 1 } 21.07.2013, 1 {, 1 } (S. & V. Korneyev) (SIZK).

Distribution. Andorra, Austria, Belgium, Britain, Bulgaria, , (mainland), , Freance (mainland), Germany, Hungary, Lithuania, (mainland), , Romania, Russia (European: North-Eastern, Central, 462 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Fig. 6. Rhagoletis spp. wings: 1 — R. meigenii; 2 — R. alternata; 3 — R. caucasica; 4 — R. berberidis; 5–6 — R. cerasi; 7 — R. cingulata.

Eastern), Slovakia, , Ukraine; Asian Russia: Northern Caucasus, Altai, Tyva, south of Far East; Kazakhstan, Kyrgyzstan, Japan. Host plants. Rosa acicularis Lindl., R. alberti Rgl., R. canina L., R. rugosa Th unb., R. spinosissima L., R. villosa (Kandybina, 1977; Smith & Bush, 1999).

Rhagoletis bagheera Richter & Kandybina, 1997 (fi gs 5,2, 7,7)

Rhagoletis bagheera Richter & Kandybina, 1997: 915.

Material. Paratypes 1 {, 1 }: Armenia: Asni, Vedi Distr., 5.08.1965, (V. Richter) (SIZK).

Distribution. Armenia, E Georgia. Host plant. Rhamnus pallasii (Rhamnaceae) (Richter & Kandybina, 1997). Remarks. Th is species can be recognized among the related Western Palaearctic species (R. batava, R. fl avigenualis, R. sp. near fl avigenualis) by the combination of black femora, smaller size, and diff erent host plant. Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 463

Fig. 7. Rhagoletis spp. wings: 1 — R. obsoleta, Myhiia; 2 — R. sp. near obsoleta, Mt. Hermon; 3 — R. completa; 4 — R. fl avicincta; 5 — R. zernyi; 6 — R. fl avigenualis; 7 — R. bagheera; 8 — R. batava.

Rhagoletis batava Hering, 1958 (fi gs 7, 8)

Rhagoletis batava Hering, 1958: 2; Rohdendorf, 1961: 183; Kandybina, 1977: 145; White & Elson-Harris, 1992: 388; Merz, 2001: 92; Norrbom et al., 1999: 200; Smit, 2010: 129.

Material. Th e Netherlands: Hompelvoet, Z.H., 10, 18.08.2000, 3 {, 3 } (B. v. Aartsen); Kyrgyzstan: on , numerous specimens; Asian Russia: Altai, numerous specimens (SIZK).

Distribution. Th e Netherlands, Switzerland (?); Spain (?); Russia: Northern Cauca- sus; Altai, Tyva; Kyrgyzstan. Host plant. Hippophae rhamnoides (Elaeagnaceae) (Kandybina, 1977). Remarks. Th is species can be recognized among the related western Palaearctic spe- cies (R. bagheera, R. fl avigenualis, R. sp. near fl avigenualis) by the combination of black femora, slightly larger size (WL = 3.5–4), and diff erent host plant (Hippophae rhamnoides rather than Rhamnus or Juniperus). We have seen no material from Switzerland or Spain, and records from these countries need verifi cation by reexamination of specimens. 464 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Rhagoletis berberidis Jermy, 1961 (fi gs 5, 3, 6, 4) Rhagoletis berberidis Jermy, 1961: 133, 136; Benedek, 1961: 45; Kandybina, 1962: 450, 1977: 159; White & Elson- Harris, 1992: 388; Merz, 1994: 108; Korneyev, 1997: 94; Norrbom et al., 1999: 200.

Material. Switzerland: Visp: Baltschieder, 46.31 N, 7.88 E, h = 640 m, 22.06.1989, 1 {, Betten-Talstation, 46.37 N, 8.08 E h = 850 m, 19.07.1991, 3 { (B. Merz); Ukraine: Kyiv: Lysa Hora, 50.40 N, 30.55 E, 16.07.1997, 6 {, 19.07.1997, 2 {, 1 } (V. Korneyev); Russia: Northern Caucasus: Karachaevo-Cherkessia: Teberda, Jamagat, 43.46 N, 41.76 E, 26.07.2013, 2 { (V. Korneyev); Kabardino-Balkaria: Nalchik, ex , 26.08.1960, 2 {, 9 } (Sheldeshova) (SIZK).

Distribution. Austria, Hungary; Slovakia, Switzerland; Ukraine; Russian Northern Caucasus; Armenia. Host plant. Berberis vulgaris L. (Berberidaceae) (Jermy, 1961; Benedek, 1961; Kandy- bina, 1977; Merz, 1994).

Rhagoletis caucasica Kandybina & Richter, 1976 (fi gs 6,3)

Rhagoletis caucasica Kandybina & Richter, 1976: 184; Kandybina, 1977: 171; Norrbom et al., 1999: 200.

Material. Russia: Northern Caucasus: Karachaevo-Cherkessia: Teberda, 43.43 N, 41.74 E, 7.07.2013, 1 {, 10.06.2013, 3 {, 3 } (S. & V. Korneyev), 18.07.2013, 1 {, 21.07.2013, 1 } (S. & V. Korneyev) (SIZK).

Distribution. Russian Northern Caucasus. Host plant. Berberis vulgaris L. (Berberidaceae) (Kandybina & Richter, 1976; Kan- dybina, 1977).

Rhagoletis cerasi (Linnaeus, 1758) (fi gs 1, 4; 5, 4; 6, 5–6)

Musca cerasi Linnaeus, 1758: 600; Trypeta cerasi: Loew, 1844: 361; Rhagoletis cerasi: Loew, 1862b: 45; Hendel, 1927: 75; Rohdendorf, 1936: 23, 1961: 179; Zaitzev, 1947: 5; Boller & Bush, 1974; Boller et al., 1976; Korneyev, 1983: 12; Kameneva & Korneyev, 1985: 68; White & Elson-Harris, 1992: 353; Merz, 1994: 109, 2001: 92; Mohamadzade & Rasoulian, 2009: 84; Smit, 2010: 130.

Material. Austria: Nickelsdorf, 47.92 N, 17.04 E, on mahaleb cherry, 4.06.2003, 4 { (V. Korneyev); Greece: Koupa NW of Skra, 41.07 N, 22.35 E, h = 700 m, swept from sweet cherry tree, 5.06.2002 (E. Kameneva, S. & V. Korneyev); Moldova: Vişniovca, 46.33 N, 28.44 E, 13.08.1988, 1 } (V. Korneyev); Switzerland: Visp: Visperterminen, 1600–1900 m, swept from Lonicera, 24 {, 11 }, Leuk, 46.33 N, 7.63 E, swept from Lonicera, 22.07.2004, 28 {, 4 } (S. & V. Korneyev); Ukraine: Donetsk: Donetsk, at light, 15.06.1992, 1 } (G. Popov); Khmelnytsk: Kamyanets-Podilsky Distr., “Tovtra Verbetska” Botanical Reserve, 14.07.1997, 2 } (O. Gumovsky), Kyiv: Kyiv, 50.467 N, 30.471 E, 1.07.2007, swept from sour cherry tree, 5 {, 4 }, Obukhiv, 50.136 N, 30.613 E, on sour cherry, 14.06.1983, (V. Korneyev); Mykolaiv: Myhiia, 48.00 N, 30.58 E, 21.06.2009, 4 {, idem, swept from Lonicera, 22.06.2009, 23 {, 12 }, idem, 23.06.2009, swept from Lonicera and Prunus, 11 {, 8 } (S. & V. Korneyev) (SIZK).

Distribution. Andorra, Austria, Belgium, Britain I., Bulgaria, Croatia, Czech Republic, Denmark (mainland), Finland, France (mainland, Corsica), Germany, Greece (mainland, Crete), Hungary, Italy (mainland), , Lithuania, Moldova, Th e Netherlands, Norway (mainland), Poland, Portugal (mainland), Romania, Sardinia, Sicily, Slovakia, Russia (European Territory: Center, East, South), Spain (mainland), Sweden, Switzerland, Ukraine; Asian Russia (Western Siberia), Georgia, Armenia, Turkey, Iran, Kazakhstan (Norther and North-Eastern). Host plants. Cherries (Rosaceae: Prunus cerasus L., P. avium L., P. serotina Ehrn., P. mahaleb L.) and honeysuckles (Caprifoliaceae: Lonicera tatarica L., L. xylosteum L.) (Hendel, 1927; White & Elson-Harris, 1992; Merz, 1994). Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 465

Rhagoletis cingulata (Loew, 1862) (fi gs 6, 7)

Trypeta cingulata Loew, 1862a: 76; Rhagoletis cingulata: Loew, 1873: 263; Bush, 1966: 473; White & Elson- Harris, 1992: 359; Norrbom et al., 1999: 201; Boller and Mani, 1994: 83; Merz & Korneyev, 2004; Smit, 2010: 130; EPPO, 2004: 8, 2010a: 4, 2010b: 5; 2013: 6; 2014: 6; Baugnée, 2006; Szeoke, 2006: 470; Bjeliš & Seljak, 2008: 14; Egatner et al. 2010: 158; EFSA, 2014: 3854; CABI, 2016; Bjeliš et al., 2016: 75. — Rhagoletis indiff erens: Merz, 1991: 56, 1994: 109 (misidentifi cation).

Material. Th e Netherlands: Burgh Zld., 51.69 N, 3.73 E, 9.08.2000, 2 {, 1 }, 17.08.2000, 1 } (B. v. Aartsen); USA: Michigan: Oceana Co., near Hart, 26.07.1989, 1 {, 1 } (J. Jenkins); Florida: Lake Co., Clermont, McPhail trap, 11.06.1992, 1 {, 1 } (J. Hunt); Orange Co., Orlando, 12.05.2005, 1 {, 1 } (J. McDermott); Lakeland, Steine trap, 4.05.1964, 1 { (R. Vild), 11.05.1964, 1{, 1 } (D. C. Phelps) (SIZK).

Distribution. Belgium, Croatia, Czech Republic, France, Germany, Hungary, Italy (mainland), the Netherlands, Poland, Slovenia, Switzerland; Canada, USA. Host plants. Prunus serotina Ehrn., P. pensylvanica L., P. virginiana L., P. cerasus L., P. avium L., and P. mahaleb L. (Rosaceae) (Smith & Bush, 1999). Remarks. Casually recorded in 1983, this species was recorded from Switzerland by Merz (1991) as “R. indiff erens”, but later was correctly identifi ed as R. cingulata (Boller & Mani, 1994; EPPO, 2004, 2010 a; Merz & Korneyev, 2004). European specimens oft en have widely darkened mid and hind femora, as well as the wing pattern typical for R. indiff erens. Available barcoding data of European specimens (J. Smit, unpublished data) fi t well to existing COI sequences for R. cingulata, rather than R. indiff erens, on GenBank. R. cingulata is a quarantine species, which is expected to spread to Ukraine.

Rhagoletis completa Cresson, 1929 (fi gs 5, 1, 7, 3)

Rhagoletis suavis ssp. completa Cresson, 1929: 412; Rhagoletis completa: Bush, 1966: 488; Merz, 1991: 56; 1994: 109; White & Elson-Harris, 1992: 359; Norrbom et al., 1999: 200; Verheggen et al., 2017: 1.

Material. USA: Washington: Asotin Co., Clarckston, ex Juglans regia, 17.10.1982, 1 {, 17.10.1983, 1 {, 2 }, 30.10.1988, exit 5.08.1989, 2 } (W. J. Turner) (det. J. Jenkins) (SIZK). No European material was available during this study.

Distribution. Austria, Croatia, France, Hungary, Germany, Slovenia, Spain, Switzerland (Verheggen et al., 2017); Canada, USA. Host plants. L., J. microcarpa Berl., J. hirsuta Manning, J. major (Torr.) Heller., J. regia L., J. californica S. Watson, J. hindii Rehd., J. regia L. (Smith & Bush, 1999). Remarks. According to Verheggen et al. (2017), invasive European populations of the walnut husk fl y initially were recorded from Switzerland and Italy (1988–1991). Th is species has subsequently established in least seven additional countries in Europe, but still has not reached the limits of its potential distribution, mainly by natural adult dissemination and adult hitchhiker behavior, and, to a lesser extent, transportation of larvae in fresh fruits. When the fl y is uncontrolled, 100% of walnut trees can be infested, and losses in walnut yields may be up to 80 %. Under phytosanitary control, the negative eff ect can be reduced to less than 10% yield loss. Th ere is a strong need for R. completa monitoring across European countries. In Ukraine, it is expected to invade the Transcarpatian Region.

Rhagoletis fl avicincta Enderlein, 1934 (fi gs 5, 5; 7, 4) Rhagoletis fl avicincta: Loew, 1873: 263 (nomen nudum); Rhagoletis fl avicincta Enderlein, 1934: 426; Rohdendorf, 1961: 184; Kandybina, 1961: 204; 1977: 143 (larvae); Korneyev & Merz, 1997: 57; Norrbom et al., 1999: 201; Mohamadzade & Rasoulian, 2009: 84. 466 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Material. Ukraine: Mykolaiv: Myhiia, 48.00 N, 30.58 E, swept from Lonicera, 21.06.2009, 6 {, 22.06.2009, 2 {, 23.06.2009, 3 {, 1 } (S. & V. Korneyev); Kyrgyzstan: numerous specimens swept from Lonicera in various localities (SIZK). Distribution. Ukraine; European Russia; Iran; Kazakhstan, Kyrgyzstan, Tadjikistan, Uzbekistan; Mongolia. Host plants. Lonicera korolkowii Stapf, L. stenantha Pojark., L. nummularifolia Jaub. and Spach (Caprifoliaceae) (Kandybina, 1977; Smith & Bush, 1999).

Rhagoletis fl avigenualis Hering, 1958 (fi gs 5, 6; 7, 6)

Rhagoletis fl avigenualis Hering, 1958: 3; Rohdendorf, 1961: 186; White & Elson-Harris, 1992: 388; Korneyev & Merz, 1997: 57; Norrbom et al., 1999: 201; Mohamadzade & Rasoulian, 2009: 84. — Rhagoletis zernyi: Rohdendorf, 1936: 5; Zaitzev, 1947: 6 (misidentifi cation).

Material. Kyrgyzstan: Ysyk-Köl Region: Terskey Alatau Mts, ex Juniperus sabina, 1 { (M. Kandybina); Kyrghyz Alatau Mts., 42.36 N, 73.87 E, 5–7.08.1998, 2 {, 1 } (E. Kameneva & V. Korneyev); Kazakhstan: Aksu- Djabagly Natural Reserve, ex Juniperus seravhanica, 5.08.1984, 17.08.1984, 15.09.1984, 5 { (Fisechko) (SIZK); Turkmenistan: Kopet-Dagh between Firyuza and [Iranian] border, 23.09.1930 (L. Bianchi) (ZISP).

Distribution. Georgia, Turkey, Iran, Turkmenistan, Kazakhstan, Kyrgyzstan, Tajikistan. Host plants. Juniper excelsa M. Bieb., J. seravschanica Kom., J. semiglobosa Rgl., J. sabina L., J. turkestanica Kom. (Cupressaceae) (Kandybina, 1977; Smith & Bush, 1999). Remarks. Th is species can be recognized among the related Western Palaearctic species (R. bagheera, R. batava, R. sp. near fl avigenualis) by the combination of yellow femora (only hind femur usually brown in basal one-third), smaller size, and diff erent host plant.

Rhagoletis sp. near fl avigenualis Hering Rhagoletis batava: Merz, 1994: 108 (misidentifi cation). — Rhagoletis fl avigenualis: Merz, 2006: 8 (misidentifi cation).

Material. Switzerland: Graubunden: Mezocco, 890 m, 1.07.1990, 1 {: (B. Merz) (MHNG); Visp: Vispetrterminen, 1300–1900 m, 21.07.2004, 1 {, 1 } (S. & V. Korneyev) (SIZK); idem, 1500 m, “Kreuz” (the cross on a hill), 21.07.2004, idem, 1400–1600 m, 18.05, 17.07, 20.07, 26.07.1993, 4 {, 3 } (B. Merz) (MHNG). Distribution. Switzerland. Host plants. Unknown; swept from Juniperus sp. Remarks. Th is species can be recognized from the related Western Palaearctic spe- cies (R. fl avigenualis, R. bagheera, R. batava) by the combination of black femora (including entire fore femur), smaller size, and diff erent host plant. It can be readily diff erentiated from R. fl avigenualis, which also feeds in juniper “berries”, by the femora black (mostly yellow, except base of hind femur in R. fl avigenualis) and by the wing crossbands uniformly dark brown (yellow, brown bordered in R. fl avigenualis). It diff ers from R. batava and R. bagheera by the fore femur largely black and the scutellum only narrowly darkened on anterior 0.1–0.2 (in R. batava and R. bagheera, fore femur only with black dorsal vitta, and scutellum widely darkened on anterior 0.25–0.30). In addition, it diff ers from R. batava by its smaller size (WL = 3.0–3.5 mm; in R. batava, WL =3.5–4.0 mm). Molecular analysis (Hulbert, Smith & S. Korneyev, in prep.) supports the hypothesis that this is an undescribed new species, which will be subject to a forthcoming paper. Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 467

Rhagoletis meigenii (Loew, 1844) (fi gs 1, 3, 6, 1) Trypeta meigenii Loew, 1844: 316; Zonosema meigenii Loew, 1862b: 44; Zonosema meigeni: Rohdendorf, 1961: 194; Kandybina, 1961: 209; Rhagoletis meigeni: Hendel, 1927: 74; Rohdendorf, 1936: 23; Kandybina, 1977: 165; Rhagoletis meigenii: Korneyev, 1983: 12; Kameneva & Korneyev, 1985: 68; White & Elson-Harris, 1992: 388; Merz, 1994: 109, 2001: 92; Korneyev, 1997: 95; Norrbom et al., 1999: 201; Merz & Korneyev, 2004.

Material. Ukraine: Kyiv: National Botanical Gardens, 50.42 N, 30.56 E, 24.07.2006, 10 } (V. Korneyev & E. Kameneva); Lysa Hora, 50.40 N, 30.55 E, 19.07.1997, 2 } (V. Korneyev); Motovylivka, 50.20 N, 30.05 E, 15.07.1985, 1 } (V. Korneyev); Cherkasy: Kaniv Distr., Trostyanets, 49.77 N, 31.36 E, 20.06.1980, 3 } (E. Kameneva); Russia: Northern Caucasus: Karachaevo-Cherkessia: Teberda, 43.43 N, 41.74 E, 10.06.2013, 4 {, 4 }, 19.07.2013, 2 }, 21.07.2013, 6 }, 22.07.2013, 4 }, 23.07.2013, 2 }, 24.07.2013, 12 }, 29.07.2013, 1 }, 30.07.2013, 1 }, 2.08.2013, 7 }, idem, Jamagat, 43.46 N, 41.76 E, 26.07.2013, 1 }, (S. & V. Korneyev).

Distribution. Spain, Latvia, ; Ukraine; Russia: European Territory, Northern Caucasus (fi rst record). Host plant. Berberis vulgaris L. (Kandybina, 1977).

Rhagoletis obsoleta Hering, 1936 (fi gs 5, 7; 7, 1) Rhagoletis cerasi f. obsoleta Hering, 1936: 182; Rhagoletis obsoleta: Rohdendorf, 1961: 182. — Rhagoletis cerasi (in synonymy): Norrbom et al., 1999: 201; Merz & Korneyev (2004) (partly).

Material. Ukraine: Mykolaiv: Myhiia, 48.00 N, 30.58 E, swept from Lonicera, 23.06.2009, 1 { (S. & V. Korneyev) (SIZK).

Host plant. Lonicera sp. (Kandybina, 1977). Remarks. Th is species was briefl y described by Hering (1936) as a form of R. cerasi without the accessory dark crossband on the wing and in a comparison with R. reducta Hering from northeastern China. Subsequently Rohdendorf used Rhagoletis obsoleta as the valid name for a single specimen from the Voronezh Region of southwestern Russia. the name became available and then used in synonymy with R. cerasi but without sound proofs. All the known records (Hering, 1936; Rodendorf, 1961; current paper) are based on single specimens collected together with numerous R. cerasi or also with R. fl avicincta on Lonicera bushes. It shows indirectly that the specimens identifi ed as R. obsoleta are merely the dark morph of R. cerasi with the accessory band completely fused with the subapical crossband. All of them have a very narrow hyaline gap between the discal and subapical crossbands (fi g. 7, 1) as if the accessory band were incorporated into the subapical cross- band, as well as entirely black femora and abdominal tergites, which is common for many European populations of R. cerasi from Lonicera. In addition, many specimens of R. cerasi have the accessory band connected or partly fused to the subapical crossband. Th e synony- my of the two names is very highly probable, but further proof based on molecular analysis of specimens from the type locality of R. obsoleta (Berlin-Frohnau) is needed. An alternative hypothesis that R. obsoleta could be a hybrid form of R. cerasi and R. fl a- vicincta, though less probable, must be also checked, as all three nominal species occur on Lonicera in southwestern Ukraine and, according to Rohdendorf (1961), in southwestern European Russia (Voronezh).

Rhagoletis sp. near obsoleta (fi gs 5, 8; 7, 2) Material. Israel: Mt Hermon, 33.38N, 35.83E, 1600 m, on Lonicera arborea, 18.07.1995, 1 {, 1 }, idem, ex berries of L. arborea, 3.08.1995 — exit 05.1996, 2 } (A. Freidberg), idem, 165o m, 17–18.07.1995, 2 { (I. Yarom) (SIZK).

Remarks. Th e specimens from Mt. Hermon precisely fi t the characters of R. obsoleta (all the femora and abdominal tergites black, wing with four wide dark brown crossbands, without traces of the accessory band). Apparently all the specimens in this locality and the 468 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev only population of a Rhagoletis species known from Israel do not vary in their wing pat- terns (unlike the European R. cerasi / R. obsoleta), but it has not been ever recorded in the literature due to its dubious species identity. It is quite possible that this geographically isolated population represents a new species. Further molecular analysis and comparison with European species is necessary.

Rhagoletis zernyi Hendel, 1927 (fi g. 7, 5) Hendel, 1927: 76; Merz, 2001: 92; Merz & Blasco-Zumeta, 1995: 132; Norrbom et al., 1999: 202. Material. Spain: Monegros, Pina-de-Negro, 13.08.1992, 1{ (Blasco-Zumeta) (Merz det. 1994) (SIZK).

Distribution. Spain. Host plants. Juniperus thurifera L. (Merz & Blasco-Zumeta, 1995).

My thanks are due to Allen L. Norrbom and anonymous referee for reviewing this manuscript and their criticism and valuable comments, and Bernhard Merz and Bernard Laundrie (MHNG) and Amnon Freidberg (TAUI), who kindly put at our disposal important material, both type and undetermined, or provided necessary data on the specimens deposited in collections under their care.

References Baugnée, J. Y. 2006. Contribution à la connaissance des Tephritidae de Belgique (Diptera: Brachycera). Notes faunistiques de Gembloux, 59, 63–113. Becker, T. 1903. Ägyptische Dipteren. (Fortsetzung und Schluss). Mitteilungen aus dem Zoologischen Museum in Berlin, 2 (3), 67–195. Bezzi, M. 1910. Restaurazione del genere Carpomyia (Rond.) A. Costa. Bollettino del Laboratorio di Zoologia Generale e Agraria della Regia Scuola d’Agricoltura, Portici (1911) 5, 3–33. Bigot, J. M. F. 1891. Th e Baluchistan melon fl y. Indian Museum Notes, 2, 51. Bjeliš, M., Buljubašić, I., Oštrkapa-Međurečan, Ž. 2016. Slow establishment of Rhagoletis cingulata in Croatia. Bulletin of Insectology, 69 (1 supplement), 75–80. Bjeliš, M., Seljak, G. 2008. Rhagoletis cingulata L. (Diptera, Tephritidae) — new quarantine pest in Croatia and Slovenia. In: 5 simpozij of plant protection in Bosnia and Herzegovina.Sarajevo, 16–18 December 2008. Zbornik rezimea, 14 Boller, E. F. & Bush, G. L. 1974. Evidence for genetic variation in populations of the European cherry fruit fl y, Rhagoletis cerasi (Diptera: Tephritidae) based on physiological parameters and hybridization experi- ments. Entomologia Experimentalis et Applicata, 17, 279–293. Boller, E. F. & Mani, E., 1994. Two North American Rhagoletis spp. in Europe. IOBC/wprs Bulletin, 17, 83. Boller, E. F., Russ, K. Vallo, V., Bush, G. L. 1976. Incompatible races of European cherry fruit fl y, Rhagoletis cerasi (Diptera: Tephritidae), their originand potential use in biological control. Entomologia Experimen- talis et Applicata, 20, 237–247. Bush, G. L. 1966. Th e , cytology and evolution of the genus Rhagoletis in North America. Bulletin of the Museum of Comparative Zoology, 134, 431–562. CABI. 2016. Rhagoletis cingulata. In: Invasive species compendium. CAB International, Wallingford, UK. [online]URL: http://www.cabi.org/isc/datasheet/47051 Costa, A. 1854. Frammenti di entomologia napoletana. Annali Scientifi ci. Giornal di Scienze Fisiche, Matemati- che, Agricoltura, Industria ec. ec. ec. (Napoli), 1, 69–91. Cresson, E. T., Jr. 1929. A revision of the North American species of fruit fl ies of the genus Rhagoletis (Diptera: Trypetidae). Transactions of the American Entomological Society (Philadelphia), 55, 401–414. Curran, C. H. 1932. New North American Diptera, with notes on others. American Museum Novitates, 526, 1–13. Effl atoun, H. C. 1924. A monograph of Egyptian Diptera, Part II., Fam. Trypaneidae. Memoires de la Societe Entomologique de Egypte, 2 (2), 1–132. Enderlein, G. 1934. Dipterologica. I. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. (1933), 416–429. EFSA. 2014. Scientifi c opinion on the pest categorisation of Rhagoletis cingulata Loew. EFSA Journal, 12 (10), 3854. Egatner, A., Zeisner, N., Hausdorf, H., Blümel, S., 2010. First record of Rhagoletis cingulata (Loew) (Dipt., Tephritidae) in Austria. Bulletin OEPP/EPPO Bulletin, 40 (1), 158–162. EPPO. 2004. Rhagoletis cingulata occurs in the Netherlands, but not R. indiff erens. EPPO Reporting Service, 6 (87), 8. EPPO. 2010a. Rhagoletis cingulata occurs in Switzerland, but not R. indiff erens. EPPO Reporting Service, 7 (127), 4–5. EPPO. 2010b. First record of Rhagoletis cingulata in Belgium. EPPO Reporting Service, 7 (128), 5. EPPO. 2013. Rhagoletis cingulata detected for the fi rst time in Languedoc-Roussillon region (FR). EPPO Reporting Service, 3 (55), 6. Th e Carpomyini Fruit Flies (Diptera, Tephritidae) of Europe, Caucasus… 469

EPPO. 2014. First report of Rhagoletis cingulata in the Czech Republic. EPPO Reporting Service, 10 (186), 6–7. Fallén, C. F. 1814. Beskrifning ofver de i sverige funna tistel-fl ugor, horande till dipter-slagtet Tephritis. Kung- liga Svenska Vetenskapakademiens Handlingar. 35, 156–177. Fallén, C. F. 1820. Ortalides sveciae. Berling, Lundae, 1–34. Freidberg, A. 2016. New taxa of Carpomyini, with special emphasis on Goniglossum (Diptera: Tephritidae: ). Zootaxa, 4144 (1), 54–70. doi: 10.11646/zootaxa. 4144.1.2. Freidberg, A., Kugler, J. 1989. Fauna Palaestina. Insecta IV. Diptera: Tephritidae. Israel Academy of Sciences and Humanities, Jerusalem: [i-vi] + 1–212, pls. 1–8, 1 map. Hendel, F. 1914. Die Gattungen der Bohrfl iegen. (Analytische Übersicht aller bisher bekannten Gattungen der Tephritinae.). Wiener Entomologische Zeitung, 33: 73–98. Hendel, F. 1927. 49. Trypetidae. In: Lindner, E. Ed. Die Fliegen der palaearktischen Region. Sweizerbart, Stutt- gart, 5 (Lfg. 16–19), 1–221. Hering, E. M. 1936. Bohrfl iegen aus der Mandschurei. (11. Beitrag zur Kenntnis der Trypetidae.). Konowia, 15, 180–189. Hering, E. M. 1958. Zwei neue paläarktische Rhagoletis (Dipt., Trypet.). (55. Beitrag zur Kenntnis der Trypeti- dae). Stuttgarter Beiträge zur Naturkunde, 7, 1–4. Hulbert, D. L., Korneyev, V. & Smith, J. J. 2016. Updating the phylogeny of Rhagoletis: focus on relationships of the North American species groups. Poster presented at International Congress of Entomology, Orlando, Florida, USA, 25–30 September 2016. Kameneva, E. P. & Korneyev, V. A. 1985. Tephritid fl ies (Diptera: Tephritidae) of the Ukrainian SSR fauna. Problems of general and molecular biology. Kiev, 4, 67–71 [In Russian]. Kandybina, M. N. 1961. On the diagnostics of the larvae of fruit fl ies (Diptera, Trypetidae). Entomologichekoe Obozrenie, 40, 202–213 [In Russian]. Kandybina, M. N. 1962. On the diagnostics of the larvae of fruit fl ies (Diptera, Trypetidae) II. Entomologichekoe Obozrenie, 41, 447–456 [In Russian]. Kandybina, M. N. 1965. On the larvae of fruit-fl ies of the genus Capomyia A. Costa (Diptera, Trypetidae). En- tomologichekoe Obozrenie, 44, 665–672. Kandybina, M. N. 1977. Larvae of fruit-infesting fruit fl ies (Diptera, Tephritidae). Nauka, Leningrad, 1–210 [In Russian]. Kandybina, M. N., Richter, V. A. 1976. A new species of the picture-winged fl y genus Rhagoletis Loew (Diptera, Tephritidae) from North Caucasus. Doklady Akademii Nauk Armyanskoy SSR, 62, 184–188. Korneyev, V. A. 1983. Tephritid fl ies (Diptera: Tephritidae) of the Middle Dnieper Territory. manuscript de- posited in the Ukrainian Institute of Scientifi c & Technical Information [UkrNIINTI], Kiev (1.12.1983, No 1343, Uk-D83), 1–28 [In Russian]. Korneyev, V. A. 1996. Reclassifi cation of Palaearctic Tephritidae (Diptera). Communication 3. Vestnik Zo- ologii, (1995) (5–6), 25–48. Korneyev, V. A. 1997. Rhagoletis berberidis (Diptera, Tephritidae, Trypetinae), the fi rst faunistic record from Ukraine. Vestnik Zoologii, 31 (5–6), 94–95. Korneyev, V. A., Dirlbek, J. Th e fruit fl ies (Diptera: Tephritidae) of Syria, Jordan and Iraq. Studia dipterologica (2000), 7 (2), 463-482. Korneyev, V. A., Merz, B. 1997. A new species of Rhagoletis Loew (Diptera: Tephritidae), with notes on Central Asian species. Journal of Ukrainian Entomological Society, 3 (1), 55–64. Korneyev, V. A. & Ovchinnikova, O. G. (2004) 79. Tephritidae — pestrokrylki [fruit fl ies]. In: Lehr, P. A., ed., Key to the of Russian Far East. Vol. VI. Diptera and Siphonaptera. Pt. 3, Dal’nauka, Vladivostok, 456–564. [In Russian] Kryshtal, O. P. 1949. [Materials to the study of entomofauna of the Middle Dnipro valley. I.] Kyiv University Publishers, Kyiv, 1–294 [In Ukrainian]. Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum cara- cteribus, diff erentiis, synonymis, locis. Tomus I. Editio decima, reformata. L. Salvii, Holmiae, 1–824. Loew, H. 1844. Kritische Untersuchung der europäischen Arten des Genus Trypeta Meig. Zeitschrift fuer Ento- mologie (Germar), 5, 312–437. Loew, H. 1856. Neue Beitrage zur Kenntniss der Dipteren. Vierter Beitrag. Programm der Koeniglichen Real- schule zu Meseritz, 1856, 1–57. Loew, H. 1862a. Diptera Americae septentrionalis indigena. Centuria secunda. Berliner Entomologische Zeits- chrift , 6, 185–232. Loew, H. 1862b. Die europäischen Bohrfl iegen (Trypetidae). W. Junk, Wien, 1–128 p. Loew, H. 1873. Monographs of the Diptera of North America. Part III. Smithsonian Miscelaneous Collection, 11 (3 [= pub. 256]): i–vii + 1–351 + I–XIII. Mamedov, D. Sh., Zhikharevich, G. P. & Mamedova, G. D. 2015. Main pests of the unabis and development of the agrotechnical measures of control them in Apsheron. Uspekhi sovremennoy nauki, (3), 53–58 [In Russian]. Meigen, J. W. 1826. Systematische Beschreibung der bekannten europaischen zweifl ugeligen Insekten. Funft er Th eil. Schulz, Hamm, i–xii + 1–412. 470 V. A. Korneyev, R. I. Mishustin, S. V. Korneyev

Merz, B. 1991. Rhagoletis completa Cresson und Rhagoletis indiff erens Curran, zwei wirtschaft lich bedeutende nordamerikanische nordamerikanische Fruchtfl iegen neu fur Europa. Mitteilungen der Schweizerischen Entomologischen Gesellschaft , 64, 55–57. Merz, B. 1994. Diptera: Tephritidae. Insecta Helvetica Fauna. HGE Press, Geneva, 10, 1–198. Merz, B. 2001. Faunistics of the Tephritidae (Diptera) of the Iberian Peninsula and the Baleares. Mitteilungen der Schweizerischen Entomologischen Gesellschaft , 74, 91–98. Merz, B. 2006. Th e 3rd Tephritoid Taxonomists’ Meeting. Instrumenta Biodiversitatis, Muséum d’histoire natu- relle, Genève, 7, 7–13. Merz, B. & Blasco-Zumeta, J. 1995. Th e fruit fl ies (Diptera, Tephritidae) of the Monegros region (Zaragoza, Spain), with the record of the host plant of Rhagoletis zernyi Hendel, 1927. Zapateri — Revista aragonesa de entomologia, 5, 127–134. Merz, B., Korneyev, V. A. 2004. Tephritidae. Fauna Europaea, version 1.1, http: //www.faunaeur.org (accessed 6.05.2016). Mohamadzade Namin, S. 2016. New distributional data on the fruit fl ies (Diptera: Tephritidae) in Iran. Ukrain- ska Entomofaunistyka, 7 (1), 47–52. Mohamadzade, S. & Rasoulian, G. R. 2009. Fruit fl ies of the genus Rhagoletis Loew (Diptera: Tephritidae) of Iran and bordering countries, with the key to species. Vestnik Zoologii, 43 (1), 81–86. Norrbom, A. L. 1997. Th e genus Carpomya Costa (Diptera: Tephritidae): new synonymy, description of the fi rst american species, and phylogenetic analysis. Proceedings of the Entomological Society of Washington, 99 (2), 338–347. Norrbom, A. L., Carroll, L. E., Th ompson, F. C., White, I. M., Freidberg, A. 1999. Systematic Database of Names. In: Th ompson, F. C., ed. Fruit Expert Identifi cation System and Systematic Information Data- base. Backhuis Publishers, Leiden, 65–299. Rekatch, V. N. 1930. Studies on biology and control of the melon-fl y Carpomyia (Myiopardalis) pardalina Zaitz. (? M. pardalina Big.). Bulletin of the Azerbaijan Central Agricultural Plant Breeding Experimental Station, 9, 1–35. Richter, V. A. 1960. New and little-known species of fruit-fl ies (Diptera, Trypetidae) in the fauna of the USSR. Entomologicheskoe Obozrenie, 39 (4), 893–896. Richter, V. A. & Kandybina, M. N. 1997. A new species of the fruit fl ies of the genus Rhagoletis (Diptera, Te- phritidae) from Transcaucaucasia. Entomologicheskoe Obozrenie, 76 (4), 914–920. Rohdendorf, B. B. 1936. [Fruit fl ies (Trypaneidae), their distribution and signifi cance as quarantine pests]. Sukhumi, NKZ Abkhazii, 1–44. Rohdendorf, B. B. 1961. Paläarktische Arten der Gattung Rhagoletis Loew (Diptera, Trypetidae) und verwandte Bohrfl iegengattungen. Entomologicheskoe Obozrenie, 40 (1), 176–213. Rondani, C. 1856. Dipterologiae Italicae prodromus. Vol. 1: Genera Italica ordinis dipterorum ordinatim dis- posita et distincta et in familiae et stirpes aggregata. Parmae, 1–228 + [1–2]. Rondani, C. 1870. Dipterologiae Italicae prodromus. Vol. 7. Genera Italica ordinis dipterorum ordinatim dispo- sita et distincta et in familias et stirpes aggregata. (Pars 4, Sect. 1). Parmae, 1–59. Silvestri, F. 1916. Sulle specie di Trypaneidae (Diptera) del genere Carpomyia dannose ai frutti di Zizyphus. Bol- lettino del Laboratorio di Zoologia Generale e Agraria della Regia Scuola Superiore d’Agricoltura, Portici, 11, 170–182. Silvestri, F. 1920. La mosca della Brionia, Gonyglossum wiedemanni Meig. (Diptera: Trypaneidae). Boll. Lab. Zool. Gen. Agrar. Portici, 14, 205–215. Smit, J. T. 2010. De Nederlandse boorvliegen (Tephritidae). Entomologische Tabellen, 5, 1–159. Smith, J. J., Bush, G. L. 1999. Phylogeny of the subtribe Carpomyina (Trypetinae), emphasizing relationships of the genu Rhagoletis. In: Aluja, M., Norrbom, A. L., eds. Fruit fl ies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, London, 187–217. Szeoke, K. 2006. First report about the occurrence of American eastern cherry fruit fl y (Rhagoletis cingulata Loew.) in Hungary. Növényvédelem, 42 (8), 470. Trikoz, N. N. & Litvinova, T. V. 2007. Damage to Ziziphus caused by the Ber Fruit Fly in the Nikita Botanical Gardens. Bulletin of the Nikita Botanical Gardens, 94, 74–76. Verheggen, F., Verhaeghe, A., Giordanengo, P., Tassus, X., Escobar-Gutiérrez, A. 2017. Walnut husk fl y, Rhago- letis completa (Diptera: Tephritidae), invades Europe: invasion potential and control strategies. Applied Entomology and Zoology, 52 (1), 1–7. White, I. M. & Elson-Harris, M. M. 1992. Fruit fl ies of economic signifi cance: their identifi cation and bionomics. International Institute of Entomology, London. i–xii + 1–601. Zaitzev, F. A. 1919. [Th e Transcaucasian melon fl y — Carpomyia (Myiopardalis) caucasica, sp. n. (Diptera, Trypetidae)]. Zapiski Nauchno-Prikladnogo Otdela Tifl isskogo Botanicheskogo Sada, 1, 64–66. Zaitzev, F. A. 1947. Th e fruit fl y fauna of the Caucasus and adjacent lands (Diptera, Trypetidae). Trudy Zoologicheskogo Instituta Akademii Nauk Gruzinskoy SSR, 7, 1–16 [In Russian].

Received 3 September 2017 Accepted 24 October 2017